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FOC | Family List | FOC Vol. 13 | Thymelaeaceae

2. Wikstroemia Endlicher, Prodr. Fl. Norfolk. 47. 1833.

荛花属 rao hua shu

Authors: Yinzheng Wang & Michael G. Gilbert

Wikstroemia dolichantha

Credit: Harvard University Herbaria

Capura Linnaeus; Diplomorpha Meisner.

Shrubs or subshrubs, occasionally small trees or rarely a herbaceous perennial (W. linoides), evergreen or deciduous. Leaves opposite or alternate. Inflorescence usually terminal and subterminal, rarely axillary, fascicled or solitary, spicate, racemose, umbelliform or capitate, sometimes in compound terminal panicles, often without involucre; peduncle long or short. Flowers bisexual or unisexual (Hawaii), 4- or 5-merous, subsessile or distinctly pedicellate; pedicel articulate. Calyx tube yellow or green, less often purplish, red or white, cylindric or tubular, sometimes slightly funnel-shaped, often caducous after anthesis, sometimes persistent; lobes 4 or 5, spreading. Petaloid appendages absent. Stamens twice as many as calyx lobes, in two series; filaments very short; anthers oblong, upper sometimes slightly exserted; connectives indistinct. Disk with 2 or 4, very rarely 1 or 5, scales, rarely joined at base by very narrow annulus, membranous. Ovary sessile, rarely shortly stipitate, usually ellipsoid, glabrous or hairy at apex, 1-loculed; style terminal, short, distinct or obscure; stigma large, capitate, globose, or discoid. Fruit a succulent berry or rather dry. Endosperm scanty or absent; cotyledons fleshy.

About 70 species: E Asia, Malaysia, Australia, and Pacific islands with a significant minor center of diversity in Hawaii where many species are dioecious; 49 species (43 endemic) in China.

The separation of Wikstroemia from the following genus, Daphne, is problematic. Features such as leaf arrangement, inflorescence type, and flower color are all clearly paralleled within the two genera and are of no diagnostic value, although they are mentioned in most keys to these genera. In most cases, the nature of the hypogynal disk is diagnostic, but even here there are species where the interpretation of the structure of the disk is difficult. For example, the distinction between the lateral ± square disk seen in several species of Daphne and species of Wikstroemia with a single large disk scale, or species of Wikstroemia in which the disk scales are connected by a narrow annulus, or species of Daphne with very deeply divided disks makes the division seem rather arbitrary, such that there must be a good case for combining these two genera, as was done by Halda (Genus Daphne. 2001). A review of generic delimitations in the family based on molecular data would be very desirable.

Wikstroemia ganpi (Siebold & Zuccarini) Maximowicz was indicated by Halda (loc. cit.: 158, as Daphne ganpi (Siebold & Zuccarini) Halda) to occur in Taiwan, but this species has not been recorded by botanists from Taiwan, and the basis of the record was not indicated.

Wikstroemia bodinieri H. Léveillé is a synonym of Alyxia schlechteri H. Léveillé in the Apocynaceae (see Fl. China 16: 162. 1995). Wikstroemia hemsleyana H. Léveillé is a synonym of Alstonia mairei H. Léveillé also in the Apocynaceae (loc. cit.: 156).


1 Calyx exterior glabrous (rarely with a few scattered hairs), often caducous after anthesis   (2)
+ Calyx tube exterior hairy, often persistent and enclosing fruit   (20)
       
2 (1) Flowers 5-merous   (3)
+ Flowers 4-merous   (11)
       
3 (2) Leaves always alternate, sometimes very close spaced and apparently whorled, blade 1-1.5 mm wide.   43 W. linearifolia
+ Leaves at least partly opposite, blade at least 3 mm wide   (4)
       
4 (3) Inflorescences clustered into a head; lateral veins 3-7 per side of midrib   (5)
+ Inflorescence a raceme or panicle, or spike; lateral veins 6-12 per side of midrib or obscure   (6)
       
5 (4) Older branches pale reddish or brownish gray; leaf blade with lateral veins 3 or 4 per side of midrib; disk apex entire or slightly retuse.   41 W. fargesii
+ Older branches dark purple-brown; leaf blade with lateral veins 5-7 per side of midrib; disk apex irregularly deeply or shallowly lobed.   42 W. fuminensis
       
6 (4) Inflorescence a solitary raceme   (7)
+ Inflorescence a panicle, lax or dense   (9)
       
7 (6) Peduncle 2-4 cm; racemes terminal and axillary.   44 W. scytophylla
+ Peduncle 0.7-1.1 cm; racemes terminal, 3-7-flowered   (8)
       
8 (7) Leaf blade 1-2 cm wide, leathery or thinly so; calyx yellow.   45 W. huidongensis
+ Leaf blade 0.3-0.8 cm wide, membranous; calyx white.   46 W. anhuiensis
       
9 (6) Panicle dense, spicate; leaves closely spaced, blade leathery, lateral veins white and prominent; calyx yellow.   47 W. lamatsoensis
+ Panicle lax; leaves not closely spaced; leaf blade papery, lateral veins not white and prominent; calyx white or yellowish green   (10)
       
10 (9) Leaf blade 1-2.2(-3.5) cm wide; calyx tube 6-8 mm.   48 W. trichotoma
+ Leaf blade 0.3-0.6 cm wide; calyx tube 12-13 mm.   49 W. leptophylla
       
11 (2) Leaves alternate, rarely subopposite   (12)
+ Leaves mostly truly opposite   (15)
       
12 (11) Inflorescence a much branched panicle.   6 W. longipaniculata
+ Inflorescence a compact head or raceme   (13)
       
13 (12) Calyx white or pink to purplish red; inflorescence bracteate.   10 W. glabra
+ Calyx yellow, yellow-green or green; inflorescence ebracteate   (14)
       
14 (13) Biennial or perennial herb or subshrub; leaf blade 1.2-2.5 × 0.3-0.8 cm.   7 W. linoides
+ Shrub, sometimes dwarf; leaf blade 3-8 × 0.7-2.8(-4) cm.   11 W. nutans
       
15 (11) Young stems pubescent or puberulous   (16)
+ Young stems glabrous   (18)
       
16 (15) Calyx tube 4-7 mm.   3 W. chui
+ Calyx tube 12-15 mm   (17)
       
17 (16) Leaf blade elliptic, ovate, or ovate-lanceolate, 4-10 × 2-3.3 cm.   4 W. hainanensis
+ Leaf blade oblong-obovate to lanceolate, 1.5-3 × 0.3-0.8 cm.   12 W. salicina
       
18 (15) Leaf blade 0.8-2.5 × 0.2-1.2 cm, ± leathery.   8 W. angustifolia
+ Leaf blade 3-20 × 1.5-9 cm, membranous to papery   (19)
       
19 (18) Leaf blade 13-20 × 4.5-9 cm; peduncle 2-6 cm; calyx lobes 5-6 mm.   2 W. liangii
+ Leaf blade 3-6(-8.5) × 1.5-2.5(-4) cm; peduncle 0.7-2 cm, slender, pendulous; calyx lobes ca. 3 mm.   11 W. nutans
       
20 (1) Flowers 5-merous   (21)
+ Flowers 4-merous   (23)
       
21 (20) Inflorescences always axillary.   38 W. paxiana
+ Inflorescences mostly terminal, often on lateral branches, sometimes also axillary   (22)
       
22 (21) Stems yellow, leaves sometimes opposite; flowers shortly pedicellate.   40 W. pilosa
+ Stems purple-red; leaves always alternate; flowers sessile.   39 W. dolichantha
       
23 (20) Flowers in terminal and short, subapical, lateral fascicles forming a leafy thyrse; hypogynal disk scale very broad, almost annular, deeply 7-10-lobed.   37 W. subcyclolepidota
+ Flowers in terminal and subterminal spikes or racemes, sometimes grouped into panicles, or axillary; hypogynal disk annular or cupular or scales 1-4, linear or oblong, apically lobed or toothed   (24)
       
24 (23) Herbaceous perennial.   7 W. linoides
+ Shrub, sometimes dwarf   (25)
       
25 (24) Leaves mostly alternate, rarely also subopposite, very rarely a few opposite   (26)
+ Leaves mostly opposite or subopposite, sometimes also alternate   (37)
       
26 (25) Inflorescence a panicle of terminal and subterminal spikes or heads; lateral leaf veins often obscure   (27)
+ Inflorescence a head elongating into a short raceme during anthesis, or a 1-4-flowered fascicle terminal on lateral branches; lateral leaf veins distinct   (29)
       
27 (26) Calyx 3.5-4 mm; leaf blade linear to narrowly oblong, 2-5 mm wide, adaxially glabrous.   32 W. cochlearifolia
+ Calyx 8-9 mm; leaf blade elliptic or obovate-elliptic or narrowly elliptic, 3-8(-15) mm wide, adaxially hairy   (28)
       
28 (27) Leaf blade adaxially tomentose; ovary densely appressed hirsute; disk scales 2, unequal, apex 2- or 3-toothed.   31 W. pampaninii
+ Leaf blade adaxially sericeous; ovary glabrous except for pilose apex; disk scale 1, apex obtuse to truncate, finely dentate or irregularly crenulate.   20 W. haoi
       
29 (26) Inflorescence a 1-4-flowered fascicle   (30)
+ Inflorescences elongating during anthesis, mostly more than 4-flowered; ovary completely pubescent; bud globose, densely white pubescent; flowers yellow   (34)
       
30 (29) Both surfaces of leaf blade glabrous, membranous   (31)
+ Both surfaces of leaf blade pubescent   (32)
       
31 (30) Leaf blade broadly elliptic to obovate, 2-5 × 1-2.5 cm; calyx pale green.   26 W. techinensis
+ Leaf blade oblong to elliptic, 0.8-1.8 × 0.4-1.1 cm; calyx yellow.   27 W. baimashanensis
       
32 (30) Leaf blade 1-2.5 mm wide.   28 W. sinoparviflora
+ Leaf blade 4-12 mm wide   (33)
       
33 (32) Leaf blade 3-3.8 × 0.9-1.2 cm.   29 W. alternifolia
+ Leaf blade 0.7-1.2 × 0.4-0.7 cm.   30 W. lungtzeensis
       
34 (29) Ovary sessile; leaf blade appressed sericeous.   33 W. capitatoracemosa
+ Ovary stipitate; leaf blade pubescent at least on midrib, sometimes glabrescent   (35)
       
35 (34) Leaf blade glabrescent, reticulate veins abaxially obscure.   34 W. ligustrina
+ Leaf blade persistently pubescent, at least on midrib, reticulate veins abaxially conspicuous   (36)
       
36 (35) Calyx ca. 15 mm, exterior with appressed and long sericeous pubescence; leaf blade 2-5 × 0.8-1.6 cm, lateral veins 5 or 6 pairs, abaxially conspicuous.   35 W. canescens
+ Calyx ca. 12 mm, exterior with appressed and short pubescence; leaf blade 1.5-3 × 0.5-1 cm, lateral veins 3 or 4 pairs, slender, abaxially obscure   36 W. lichiangensis
       
37 (25) Leaf blade 1.5-2.5 mm wide, to 2.5 cm.   23 W. stenophylla
+ Leaf blade (2-)3-35 mm wide, if less than 3 mm, then at least 2.5 cm   (38)
       
38 (37) Calyx pink, purple, lilac, or lavender   (39)
+ Calyx yellow, greenish yellow, yellowish white, or green, sometimes flushed with purple or red   (42)
       
39 (38) Calyx 4-6 mm   (40)
+ Calyx 7-12 mm   (41)
       
40 (39) Ovary glabrous except for pilose apex; inflorescence ± paniculate, ca. 20-flowered.   20 W. haoi
+ Ovary uniformly hairy; inflorescence not paniculate, 3-5(-8)-flowered.   21 W. vaccinium
       
41 (39) Leaf blade drying dark brown; lateral veins 4 or 5 per side; calyx tube 9-12 mm.   17 W. monnula
+ Leaf blade drying green, lateral veins 10 or 11 per side; calyx tube ca. 15 mm.   1 W. mononectaria
       
42 (38) Calyx 4-6 mm   (43)
+ Calyx 7-15 mm   (44)
       
43 (42) Calyx yellowish white or green, sometimes flushed with purple; tube slightly ovoid, 2-5 mm.   15 W. taiwanensis
+ Calyx yellow; tube cylindric, 4-6 mm.   9 W. micrantha
       
44 (42) Inflorescence a panicle of racemes   (45)
+ Inflorescence capitate or shortly racemose, sometimes indistinctly paniculate   (48)
       
45 (44) Leaf blade abaxially glabrous; calyx greenish yellow or creamy-yellow, sometimes flushed with purple, tube often less than 10 mm   (46)
+ Leaf blade abaxially hairy; calyx yellow, tube ca. 10 mm   (47)
       
46 (45) Leaf blade lanceolate-obovate, obovate, or oblanceolate, 1.6-2.5 cm wide; hypogynal disk annular, extended on one side into 1(-3) obovate scale(s).   5 W. delavayi
+ Leaf blade lanceolate to oblong-lanceolate, 0.2-1 cm wide; hypogynal disk scales 2(or 3), linear or oblong, basally connate.   22 W. chamaedaphne
       
47 (45) Leaf blade elliptic or ovate-orbicular to oblong, 0.8-2.8 cm wide; hypogynal disk scales 2, one linear and one oblong.   24 W. gracilis
+ Leaf blade narrowly oblanceolate to broadly linear, 0.4-0.8 cm wide; hypogynal disk scale 1, obovate, margin crenate.   25 W. reginaldi-farreri
       
48 (44) Lateral veins slender, dense, at a very narrow angle to midrib   (49)
+ Lateral veins remote, obscure, 5-7 pairs, at wide angle to midrib   (50)
       
49 (48) Leaf blade oblong to lanceolate, 5-15 mm wide, abaxially reddish brown when dried.   14 W. indica
+ Leaf blade elliptic or oblong, apex broad, 4-9 mm wide, both surfaces glabrous, abaxially pale green.   18 W. capitata
       
50 (48) Leaf blade lanceolate, 0.8-1.2 cm wide; stems relatively slender.   16 W. lanceolata
+ Leaf blade obovate to oblong, 1.3-2.5 cm wide; stems relatively stout   (51)
       
51 (50) Inflorescence often 15-26-flowered, peduncle to 0.5 cm, stout.   13 W. pachyrachis
+ Inflorescence 4-6(-20)-flowered, peduncle 0.7-3 cm, slender.   19 W. retusa

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